Special Issue Contemp.Trends.Geosci.,7(3),2018,255-282 DOI: 10.2478/ctg-2018-0018

Late ‘odds and ends’ from the Low Countries

John W.M. Jagt1,*, Barry W.M. van Bakel2, Mart J.M. Deckers3, Stephen K. Donovan4, René H.B. Fraaije2, Elena A. Jagt-Yazykova5, Johan Laffineur6, Eric Nieuwenhuis7, Bruno Thijs8

1Natuurhistorisch Museum , de Bosquetplein 6-7, 6211 KJ Maastricht, the 2Oertijdmuseum, Bosscheweg 80, 5283 WB Boxtel, the Netherlands 3Industriestraat 21, 5931 PG Tegelen, the Netherlands 4Naturalis Biodiversity Center, and Systematics Group, Postbus 9517, 2300 RA Leiden, the Netherlands 5Uniwersytet Opolski, Katedra Biosystematyki, Pracownia Paleobiologii i Ewolucji, ul. Oleska 22, 45-052 Opole, Poland 6Olympialaan 1, bus 4, 3630 Maasmechelen, 7Hub. Ortmansstraat 4, 6286 EA Partij-Wittem, the Netherlands 8Universiteit Utrecht, Faculteit Geowetenschappen, Princetonlaan 8a, 3584 CB Utrecht, the Netherlands

*Corresponding author: [email protected]

Received: 19th July, 2018 Accepted: 29th October, 2018

Abstract From various levels within the Gulpen and Maastricht formations (upper lower to upper upper , c. 69.5–66 Ma) in the extended type area of the Maastrichtian (southeast Netherlands, northeast Belgium and the Aachen area in ), a few recent additions to echinoderm faunas are illustrated and briefly discussed. Added are some erratic, flint-preserved, pre-Maastrichtian echinoid taxa from Pleistocene fluvial gravel deposits; these are of palaeogeographical interest. Crinoids include the comatulid Semiometra saskiae with traces of sublethal predation and several bourgueticrinines. Amongst the latter, the species Dunnicrinus aequalis is found preserved in biocalcarenites as well as in flint, comprising a near-complete specimen with a spectacular, semi- closed crown and a number of dissolved thecae in nodules with flint-filled, 3D-preserved nerve canals. In addition, two types of bourgueticrinid holdfasts on echinoid tests are described. Amongst echinoids, cidaroid taxa such as Temnocidaris (T.) sp. 1, with preserved primary and scrobicular spines, and Temnocidaris (Stereocidaris) aff. arnaudi, are briefly discussed. Of special note is a near-complete, flattened test of Palaeodiadema (in a private collection) from the upper Maastricht Formation at Eben Emael (province of Liège, northeast Belgium); although no spines are associated, this specimen could well become an eye opener in diadematoid classification. Of striatoradiatus, a number of tests with growth deformities (i.e., lack of one of the paired ambulacra, healed puncture wounds and apical depressions) and syn-vivo parasite infestation have been collected over recent years. Bite marks and healed puncture wounds on tests of Echinocorys spp. are recorded from the Vijlen and Lixhe 1 members (Gulpen Formation), in addition to bioerosional trace fossils produced by acrothoracican cirripedes and anomiid bivalves. Clusters of phymosomatid, holasteroid and spatangoid echinoids testify to obrution-induced mass mortality, possibly following(?)spawning. Added is the first record from the Maastrichtian type area of echinoid-produced traces (comparable to Scolicia), based on a slab from the basal Valkenburg Member; associated spatangoids comprise only members of the micrasterid genus Diplodetus. Of ophiuroids, a fairly rich material of Ophiomusium, or a closely related genus, is available from the Gronsveld, Emael and Meerssen members,

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preserving (partial) discs with arms attached that reveal all details of disc plating, mouth frame and arm structure. Asteroids include a (?)new astropectinid from the uppermost Gronsveld Member that appears to be distinct from Betelgeusia exposita, Aldebarania taberna and other paxillosidans on record from the area. This is preserved in life position, directly below a mass accumulation of both marine (sea grass) and terrestrial plants that testifies to a sudden event (obrution). A near-complete specimen of the goniasterid Chomataster acules, contained in a flint nodule, adds to our knowledge of this taxon by preliminary CT scanning. Pre-Maastrichtian, flint-preserved erratics include a test of Bathysalenia and isolated primary spines of Hirudocidaris. The former genus is known from the area, from lower upper Campanian strata (B. obnupta), but the apical disc and ambulacral plating of the present specimen appears to differ from that of both older (Turonian-Coniacian) populations of B. granulosa and the younger B. obnupta. The genus Hirudocidaris is unknown from the study area and, therefore, is assumed to have come from further afield (northwest France, southern Belgium)..

Key words: Echinoidea, Ophiuroidea, Asteroidea, Crinoidea, erratic boulders, northwest

Introduction reference is made to Jagt (1999a) and Jagt and Jagt-Yazykova (2012) (Fig. 2A, B). Below we Right from the start of serious fossil collecting present recent additions to echinoderm faunas in the Maastricht area in the latter half of the that are of definite Maastrichtian date, having eighteenth century, (echinoids in been collected from the Vijlen (intervals 5 and particular) have been attracting attention 6), Lixhe 1-3 and Lanaye members (Gulpen (Leske, 1778; Faujas de Saint-Fond, 1798- Formation) and from all members of the 1803). During the two centuries that followed, overlying Maastricht Formation. A small numerous species of echinoid, asteroid, crinoid number of specimens have been taken from and ophiuroid have been described in scientific existing museum collections, primarily from the and popular literature alike, and their Natuurhistorisch Museum Maastricht (NHMM; stratigraphical ranges refined. Dissociated EN – Eric Nieuwenhuis Collection; JJ – John ossicles of holothurians have also been Jagt Collection; LJ – Johan Laffineur recorded, albeit rarely (Zeleznik, 1985; Reich, Collection; LN – Leo Nelissen Collection; MD 2003); more research in this field is needed. – Mart Deckers Collection; MK – Werner M. Here we present brief descriptions of a Felder Collection) and the Oertijdmuseum, selection of newly collected echinoderms, as Boxtel (MAB). well as softground and bioerosional ichnofossils Over recent decades, non-professional linked with these. These specimens originate palaeontologists, from all walks of life (Fig. 2C, from the middle and upper Gulpen Formation D), have been instrumental in furthering our (Vijlen, Lixhe and Lanaye members) and entire knowledge of faunal and floral Maastricht Formation (Valkenburg, Gronsveld, assemblages in the extended area of the Schiepersberg, Emael, Nekum and Meerssen Maastrichtian Stage. This paper is a tribute to members) as exposed at a number of localities all of them. in the extended type area of the Maastrichtian Stage (Fig. 1). Brief descriptions, arranged by group

Study area and stratigraphy Crinoids

For recent overviews of lithostratigraphical and Locally, comatulid crinoids of the genera biostratigraphical subdivisions of Upper Amphorometra Gislén, 1924, Jaekelometra Cretaceous strata in the study area (Fig. 1), Gislén, 1924 and Semiometra Gislén, 1924 are

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Fig.1. Map of the extended type area of the Maastrichtian Stage, showing key localities in southern (the Netherlands, N), in the provinces of Liège and Limburg (Belgium, B) and in the Aachen area (Germany, G) [inset]. common in the Nekum and Meerssen members indentations in the margin that are here of the Maastricht Formation (Jagt, 1990, interpreted as bite/pinch marks of some 1999b). Centrodorsals, occasionally with the predator, either a fish or a decapod crustacean. basal and radial rings still articulated and Other protential predators on such comatulids illustrating various ontogenetic stages, are include ammonoid, nautiloid and coleoid sturdy and easily recognised. Associated are cephalopods and regular echinoids. Stevenson both proximal and distal brachials and cirrals, et al. (2017) have recently documented that which suggests some site fidelity and relatively even regular echinoids (e.g., cidaroids) can feed limited post-mortem transport. The small, on co-occurring comatulids. This may also have sturdy and conical centrodorsals of been the case in the upper Maastricht Formation Jaekelometra spp. are either of (near-) pristine from which a number of cidaroid taxa have been preservation, or show signs of fracturing and recorded (Jagt, 2000b; see below). abrasion, with all intermediates. Fragmentary Of stalked crinoids, additional material of specimens may be illustrative of predation and Dunnicrinus aequalis (d’Orbigny, 1841) has selective cropping, passing through the stomach been collected, subsequent to detailed accounts and intestines of fish (or other larger of a crinoid/asteroid/ophiuroid Fossil predators/scavengers) and being partially Lagerstätte from the lower Gronsveld Member abraded in the process. Others may be truly (St. Pieter and ENCI horizons, Maastricht sublethal. The average-sized centrodorsal of Formation) at the ENCI-HeidelbergCement Semiometra saskiae Jagt, 1999b illustrated here Group quarry (Jagt et al., 1998; Jagt, 1999b). (Fig. 3A-C) shows two conspicuous Specimens originating from biocalcarenites

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al., 2012; Donovan and Jagt, 2014) (Fig. 6A-C). Two distinct types of holdfasts, 1 and 2, can be distinguished. Type 1 has numerous ‘roots’, of variable length and thickness and with distinct external sutures between ankylosed radicular elements, while type 2 generally is smaller and more disc like with short marginal extensions. In addition, this type often occurs in clusters (Fig. 6C). It would seem that, in the absence of the associated stalk and aboral cups, these cannot be identified to species. In the Vijlen and Lixhe 1-2 members, the commonest bourgueticrinid species with a figure-8-shaped lumen is Bourgueticrinus constrictus (von Hagenow, in Quenstedt, 1876).

Fig.3A-C. Semiometra saskiae Jagt, 1999b, NHMM EN 83s, centrodorsal (greatest diameter: 9 mm) with sublethal predation traces (arrows) in ventral, dorsal and lateral views, ‘t Rooth quarry, Bemelen; lower Meerssen Member, Maastricht Formation (photograph: Eric Nieuwenhuis). include a near-complete individual, with a spectacular, semi-closed crown (Fig. 4A, B). This corroborates an earlier interpretation of the number of syzygial brachial articulations in this Fig.4. Dunnicrinus aequalis (d’Orbigny, 1841), species (Jagt, 1999b). In addition, we have a MAB 005514 (leg. Aad van den Engel), near- number of dissolved thecae in nodules with complete specimen (A; greatest length: 273 mm) flint-filled, 3D-preserved nerve canals (Fig. 5A- with a spectacular, semi-closed crown (close-up, B; C). greatest height, including proximale: 99 mm), part of There are also additional examples of the stalk missing (arrow); ENCI-HeidelbergCement bourgueticrinid holdfasts on medium- and Group quarry (Maastricht); basal Gronsveld Member, Maastricht Formation (photographs: Mart large-sized echinoids of the genus Echinocorys Deckers). Leske, 1778, adding to previous finds (Jagt et

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formally named and, with the exception of those of certain psychocidarids (Tylocidaris Pomel, 1883 [sensu stricto] and T. (Oedematocidaris) Smith and Wright, 1989) and the genus Hirudocidaris Smith and Wright, 1989, that such spines cannot be identified to species unless associated with (partial) tests.

Fig.5. Dunnicrinus aequalis (d’Orbigny, 1841); A, C. Dissolved thecae (LJ 025a [diameter: 2 mm] and LJ 302 [greatest height: 8 mm], respectively) in nodules with flint-filled, 3D-preserved nerve canals; residual flint deposits at CBR-Lixhe quarry (province of Liège, Belgium); B. Dissolved theca and proximal columnals [L. Nelissen Collection, unregistered; greatest height 4 mm] from Pleistocene River Maas gravel deposits (photographs: Johan Laffineur [A, C] and Mart Deckers [B]).

Echinoids

Amongst echinoids, the cidaroid Temnocidaris (T.) sp. 1 (sensu Jagt, 2000b), with preserved primary, scrobicular and ambulacral spines, is of importance. Features of both test and spines (Fig. 7A-D) differentiate it from associated congeners; moreover, this type of ambital interambulacral primary spines can now be Fig.6. Bourgueticrinid holdfasts – examples of two firmly linked to this type of test. Previously, this types (A, B: type 1; C: type 2) on medium- and large- spine type was associated with various cidarid sized echinoids of the genus Echinocorys Leske, species, as examples in Smiser (1935, pl. 1, fig. 1778; Lixhe 1 Member, Gulpen Formation; Kreco quarry (= CPL-Haccourt), Haccourt (province of 11a, as Dorocidaris faujasi; pl. 1, fig. 12a, as Liège, Belgium); A. NHMM EN 72h [greatest Typocidaris serrata; pl. 2, fig. 2a, as diameter: 4.5 mm]; B. NHMM EN 76r [greatest Stereocidaris sceptrifera) illustrate. We diameter: 16 mm]; C. NHMM EN 64v [greatest subscribe to the view expressed by Smith and diameter: 2-4 mm] (photographs: Eric Wright (1989) and Smith and Jeffery (2000) Nieuwenhuis). that isolated cidaroid spines should not be

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Fig.7. Temnocidaris (T.) sp. 1 (sensu Jagt, 2000b), MAB 005821, with preserved primary and scrobicular interambulacral spines and ambulacral miliary spines, in various aspects; CBR-Romontbos quarry, Eben Emael (Liège, Belgium); upper Nekum Member, Maastricht Formation. Greatest diameter of test: 78 mm (photograph: Mart Deckers).

Another cidaroid (Fig. 8), a species of Belgium). Although no spines are associated Temnocidaris (Stereocidaris) Pomel, 1883, (Fig. 9), this is closely comparable to appears closely related to T. (S.) arnaudi dissociated test material from the same area, as (Lambert, 1909) (sensu Smith and Jeffery, documented previously (Jagt, 2000b). A 2000). One-fifth segment of test is available revision of Late Cretaceous and early Paleogene from lower Meerssen Member at the ‘t Rooth diadematoids is under way (John W.M. Jagt, quarry, Bemelen; this is of moderate size (test Christian Neumann and Søren B. Andersen; height 38 mm), has 7 plates in an compare Smith and Wright, 1990). interambulacral column, with faint pitting along Bite traces (tooth marks), probably mostly sutures but no neural ridges. Ambital inflicted by teleost fish and/or neoselachians ambulacral plates have one large and up to 4 and comparable to examples from Upper smaller tubercles; adoral plates have one large Cretaceous levels in northern Germany marginal, a smaller median and a large perradial (Schormann, 1987; Frerichs, 2012; Girod and tubercle. Rösner, 2013) and elsewhere (Dortangs, 1998), Of special note is a near-complete, flattened occur in particular in species of the genus test of a diadematid, provisionally assigned to Echinocorys from the Vijlen Member. Being Palaeodiadema Pomel, 1883, from the upper benthic islands on a ‘soupy’ sea floor, these Maastricht Formation (?Emael or Nekum holasteroids possibly were the easy prey (Fig. members) at Eben Emael (Liège, northeast 10A-D). Corresponding tooth marks occur on

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opposite sides of the test, at the ambitus (i.e., the rather constitute traces of anchorage greatest diameter), and testify to handling by (attachment; superficial embedment) and there predators that tried to lift these tests from the sea are apically concentrated pits, some even floor by holding them between the teeth of the coalesced, of the type described recently by upper and lower jaws. At times, more than one Hammond and Donovan (2017). attempt can be postulated from partially overlapping marks (e.g., Fig. 10D). Bite marks on tests of Echinocorys from the Maastrichtian of northern Germany, as described by Thies (1985), differ in being more circular, situated more closely to the apex of the echinoid tests and in close-set clusters. Wilson et al. (2014) recorded closely comparable bite marks from Middle echinoid spines. In any case, these examples illustrate aborted attempts to feed on these echinoids; the test was merely ‘scratched’, but not pierced, unlike other examples listed below.

Fig. 9. Palaeodiadema sp. (private collection of V. Fiume, photographed at a fossil fair several years ago); CBR-Romontbos quarry, Eben Emael (Liège, Belgium); Maastricht Formation, exact level unknown, but probably Emael or Nekum members. Original diameter size about 100 mm (photograph: Stijn Goolaerts).

In addition to sublethal predation, healed scars and possible attachment sites, there are bioerosional traces of episkeletozoans such as acrothoracican cirripedes of the Rogerella de Saint-Seine, 1951 type that preferentially used ambulacral pores (Fig. 12A), as previously documented for other holasteroid genera such as Cardiaster Forbes, 1850 (Donovan et al.,

Fig.8. Temnocidaris (Stereocidaris) aff. arnaudi 2016b) and Hemipneustes Agassiz, 1836 (Lambert, 1909) [sensu Smith and Jeffery, 2000], (Donovan and Jagt, 2013). Another type can be NHMM EN 55j, one-fifth segment of test (greatest linked to anomiid bivalves: Centrichnus test height: 38 mm); ‘t Rooth quarry, Bemelen; lower eccentricus Bromley and Martinell, 1991 (Fig. Meerssen Member, Maastricht Formation 12B). To date, the latter ichnofossil has only (photograph: John W.M. Jagt). been described from echinocorythids, which is

Healed injuries, of various shapes and sizes, a conundrum since various other medium- to and either repaired or not (Fig. 11A-D); some of large-sized holasteroids are present in the Gulpen and Maastricht formations and isolated these have a pentagonal shape (Donovan et al., 2008, 2010). Some of these instances may

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shells of anomiids are found associated as the one described recently by Donovan et al. (compare Neumann et al., 2015). (2018). Similar to species of Echinocorys, such Of Hemipneustes striatoradiatus (Leske, medium- to large-sized were ideal benthic 1778), a number of tests with growth islands for several generations of deformities (i.e., lack of one of the paired episkeletozoan taxa (Nebelsick et al., 2007; ambulacra, a disjunct apical disc and apical Belaústegui et al., 2017). depressions) (Fig. 13A-E) and syn-vivo parasite Apical depressions in H. striatoradiatus infestation have been collected in recent years. (which are altogether rare) vary widely in depth Others have large circular holes (Fig. 14) with a (Fig. 15A, B); in some the apical disc is missing. different upper and lower diameter, testifying to There are comparable examples in much post-mortem boring by certain species of smaller-sized spatangoids such as Cyclaster gastrochaenid and/or mytilid [lithophagine] danicus (Schlüter, 1897) (see Asgaard, 1976), bivalve, leading to the trace fossil Bolbaster prunella (Lamarck, 1816) (see Jagt, Gastrochaenolites Leymerie, 1842 (Fig. 14). As 2000b, pl. 29, fig. 6) and species of Echinocorys far as syn-vivo infestation is concerned, most (see Schneider, 2018). Apparently, infestation examples can be ascribed to Oichnus excavatus occurred early during ontogeny; all examples Donovan and Jagt, 2002 (= Tremichnus currently available are in ‘normal-sized’ tests, excavatus; sensu Wisshak et al., 2015), with suggesting that growth was not greatly rare occurrences of heavily infested tests such hindered.

Fig. 10. Predatory traces (tooth marks; average length 5-10 mm) on Echinocorys gr. limburgica Lambert, 1903; A, B. NHMM EN 51k; C. NHMM EN 58y; D. NHMM EN 64g; Kreco quarry (= CPL-Haccourt), Haccourt (Liège, Belgium); Vijlen Member (intervals 5-6), Gulpen Formation (photographs: Eric Nieuwenhuis).

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Fig. 11. Examples of sublethal injuries and scalloped attachment scars in the test of Echinocorys gr. conoidea (Goldfuss, 1829) [sensu Jagt, 2000b). A. NHMM EN 69b [note regenerated tubercles and miliaries, but lack of pore regeneration; greatest width of scar: 24 mm]; B. NHMM EN 68w [greatest width of scar: 8 mm]; C, D. NHMM EN 68n [greatest width: 3-6 mm]; Kreco quarry (= CPL-Haccourt), Haccourt (Liège, Belgium); Lixhe 1 Member, Gulpen Formation (photographs: Eric Nieuwenhuis).

Clusters of specimens of the spatangoids within the Lanaye Member (Gulpen Formation) Diplodetus duponti (Lambert, 1911) (Fig. 16), of the Haccourt-Lixhe area (Belgium). Leymeriaster maestrichtensis (Schlüter, 1897) Also of interest are associations of small (Fig. 17) and of >50, small-sized (several size classes, with test widths ranging phymosomatids of the genus Gauthieria from 6 to 14 mm) of Cardiaster gr. granulosus Lambert, 1888 (Fig. 18), possibly (Goldfuss, 1829) (Fig. 19A, B). The smallest corresponding to spawning events and tests in this lot are comparable in size to subsequent current-induced concentrations of associated tests of Procassidulus lapiscancri (denuded) tests. Of note in the last-named (Leske, 1778) and Nucleopygus scrobiculatus example is that tests retain primary spines in a (Goldfuss, 1829). In view of the fact that slight ‘relaxed position’, suggesting that obrution may crushing (sediment loading) occurred and that be responsible for this ‘frozen behaviour’. Quite epitaxial cement tends to obscure details of pore a lot of specimens in these clusters preserve and apical disc structure, it cannot be complete apical disc plating (although details determined beyond doubt which of these size are difficult to make out) and lanterns, which is classes was truly juvenile, i.e., lacked functional indicative of limited post-mortem disturbance. gonopores. Van der Ham et al. (2006) described Non-bioerosional trace fossils (i.e., comparable examples of ‘populations’ of burrows) that could be ascribed to echinoid cassiduloids and spatangoids from flint levels bioturbation (Fig. 20) are the first to be recorded

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Fig. 12. Acrothoracican (A, Rogerella isp.; NHMM EN 68w) and anomiid bioerosional traces (B, Centrichnus eccentricus Bromley and Martinell, 1991; NHMM EN 69e, around 4 mm in greatest length) in tests of Echinocorys gr. limburgica Lambert, 1903, Kreco quarry (= CPL-Haccourt), Haccourt (Liège, Belgium); Gulpen Formation, Vijlen Member (interval 6) (photographs: Eric Nieuwenhuis). from the type Maastrichtian; these possibly tissue; successful predation would only have belonging to Scolicia. The only irregular resulted in shattered tests that could rapidly echinoids able to construct such burrow type become dispersed on the sea floor with little or known to date from this part of the sequence are no clue as to the mode of occurrence and representative of the micrasterid genus preservation. Diplodetus Schlüter, 1900. In contrast, the bioerosional trace fossil Gnathichnus pentax Ophiuroids Bromley, 1975 is well known throughout the Of ophiuroids, a rich material of Ophiomusium, sequence, from carbonate substrates, which is or a closely related genus, is available from the not surprising when the wealth of cidaroid, Gronsveld, Emael and Meerssen members, salenioid and phymosomatoid echinoids in documenting a number of species, preserving these strata is considered. (partial) discs with arms attached that reveal all bite marks have not been details of disc plating, mouth frame and arm recognised as such, despite the common structure (Fig. 21A, B). There are also rare finds occurrence and relatively high diversity of this of smothered individuals, ‘frozen’ in the group of reptiles in the area (compare Neumann process of escaping from the sediment, and and Hampe, 2018). Healed puncture wounds closely comparable to cases documented in (see above) show the resilience of stereom detail by Ishida and Fujita (2001). These pertain

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Fig.13. Hemipneustes striatoradiatus (Leske, 1778), tests with growth deformities, all from the Nekum Member (Maastricht Formation) at various localities in southern Limburg; A. NHMM MK 860A2, with extended ambulacrum I and widened test, in several aspects; B. NHMM MK 890A8, with severe deformation following a bite (?) in ambulacrum IV, in several aspects; C, D. NHMM MD 5422a, lack of ambulacrum V; E. NHMM 2015 004, lack of ambulacrum IV. Greatest test lengths between 85 and 100 mm (photographs: Mart Deckers). to Felderophiura vanderhami Jagt, 1991 (Jagt incorporated in a flint nodule, even preserves et al., 2015a) from the upper Meerssen Member knob-like spines on superomarginals. Low- and ‘Stegophiura’ hagenowi (Wienberg resolution CT images prove that the disc and all Rasmussen, 1950) from the basal Gronsveld arms are preserved (Fig. 23A, B). CT images of Member. higher resolution will be prepared to allow 3D printing of this prime specimen to allow the full Asteroids ossicular complement of this goniasterid to be described in detail. Asteroids include a (?) new astropectinid (uppermost Gronsveld Member; Fig. 22) that Echinoderms from erratic boulders, in appears to be distinct from Betelgeusia exposita particular echinoids Blake and Jagt, 2005 and Aldebarania taberna Blake and Jagt, 2005 and other paxillosidans in Over an extensive portion of the territory of the these assemblages (Jagt, 2000c). This is Netherlands, i.e., roughly the part north of the preserved in life position (with upturned arm larger rivers in the centre of the country, tips), directly below a mass accumulation of Pleistocene glaciers and the Eridanos Stream both marine (sea grass; compare Van der Ham left crystalline and sedimentary rocks of a et al., 2017) and terrestrial plants that testifies to northerly and northeasterly origin (Akkerman, a sudden event (storm). 2012). In the southeast of the country, rivers An individual of the goniasterid such as the Rhine and Maas (Meuse) added to Chomataster acules Spencer, 1913, the diversity of erratic rocks and fossils, in

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Fig.14. Circular hole, testifying to boring by certain species of bivalve (gastrochaenid or mytilid [lithophagine]), producing the trace fossil Gastrochaenolites Leymerie, 1842; CBR-Romontbos quarry, Eben Emael (province of Liège, Belgium); upper Emael Member, Maastricht Formation (J. Snellings Collection, unregistered). Greatest test length: 97 mm (photograph: Mart Deckers). delivering material from southerly areas in In the south of the country, the upper northern France, Germany and areas as far as portion of the Gulpen Formation (upper Vijlen, south as Switzerland (Bosch, 1992) that ranges Lixhe 1-3 and Lanaye members) and the in age from to Late Jurassic (Jagt Gronsveld to lower Nekum members of the et al., 2015b; Donovan et al., 2016a). overlying Maastricht Formation are Within assemblages from the north of the characterised by the regular occurrence of flint Netherlands, Van der Lijn (1974) mentioned (nodules, tabular, tubular and concretionary; irregular echinoids (mostly Galerites and Felder and Bosch, 1998a, b). A large portion of Echinocorys) and isocrinid crinoids as glacially these flint-bearing biocalcarenites was eroded transported erratic material of a northerly during the Pleistocene by the River Maas provenance. To the same category belong flint- (Meuse), which explains why flint nodules preserved (either as loose internal moulds or (mostly angular) are common in gravel deposits external moulds in cobbles and boulders) across southern Limburg (the Netherlands) and echinoids that are occasionally found on adjoining Belgian and German territories. In beaches in the north of the country or on the principle, all species listed by Van der Ham et Wadden Islands (https://historiek.net/meisje-7- al. (2006) from in-situ occurrences on vindt-fossiele-zee-egel-op-strand- Montagne Saint Pierre (Lixhe-Haccourt area, texel/16981/). province of Liège, Belgium) can be found in such boulders and cobbles. In view of the fact

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Fig.15. Hemipneustes striatoradiatus (Leske, 1788), tests with apical depressions of variable depth; ‘t Rooth quarry, Bemelen; Nekum Member (Maastricht Formation); A. NHMM 2018 009 [greatest test length: 100 mm]; B. J. Severijns Collection, no. 601; greatest test length: 84 mm; depth of depression: c. 19 mm (photographs: John Stroucken).

Fig.16. Diplodetus duponti (Lambert, 1911), NHMM EN 78n; ENCI-HeidelbergCement Group quarry (Maastricht); basal Gronsveld Member, Maastricht Formation. Greatest test width of largest specimen: 44 mm (photograph: Eric Nieuwenhuis).

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Fig.17. Leymeriaster maestrichtensis (Schlüter, 1897), NHMM MD 4561; ENCI-HeidelbergCement Group quarry (Maastricht); upper Nekum Member, Maastricht Formation. Greatest length of slab: 350 mm (photograph: Mart Deckers).

Fig.18. Gauthieria gr. radiata (Sorignet, 1850), NHMM MD 4821.07 (greatest test diameter: 11 mm); ENCI- HeidelbergCement Group quarry (Maastricht); uppermost Meerssen Member, Maastricht Formation (photograph: Mart Deckers).

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Fig.19A, B. Associations of small (several size classes; greatest test widths between 6 and 14 mm) of Cardiaster gr. granulosus (Goldfuss, 1829) and a single specimen of Procassidulus lapiscancri (Leske, 1778) in B (top centre), NHMM 2018 010a/b (leg. R. Pieters); CBR-Romontbos quarry, Eben Emael (province of Liège, Belgium); upper Nekum Member, Maastricht Formation (photographs: John Stroucken).

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Fig.20. Scolicia(?) isp., NHMM 2015 012; ENCI-HeidelbergCement Group quarry (Maastricht); basal Valkenburg Member, Maastricht Formation. Greatest length of slab: 422 mm (photograph: John Stroucken). that the stratigraphical range of the majority of appear to differ both from older (Turonian- these species is well known, these finds are Coniacian) populations of B. granulosa easily interpreted as local or regional erratic (Woodward, 1856) and of the younger B. elements. obnupta. The other comprises isolated primary However, Pleistocene gravel deposits of the spines of the cidarid Hirudocidaris of the group River Maas (Meuse) occasionally also yield of hirudo (Sorignet, 1850) (Fig. 26A, B). flint-preserved echinoid taxa that are not known Representatives of this genus are unknown from to occur in Upper Cretaceous rocks in the same Upper Cretaceous strata in the study area and area. Here we list two pre-Maastrichtian, flint- must have come from further afield (northwest preserved echinoid taxa. One of these is the France, southern Belgium). internal mould and an associated partial The well-rounded, small-sized 'Maaseieren' external imprint of the same test of a species of (Meuse or flint eggs) also contain other Bathysalenia (Figs. 24A-C, 25A-E). Although macrofossil taxa, amongst which are thick- this genus is known from the area, from lower shelled inoceramid bivalves that are distinct upper Campanian strata [B. obnupta (Schlüter, from indigenous species in Upper Cretaceous 1892); see Jagt, 2000b], both the apical disc and (lower Campanian-lower Maastrichtian) strata in the study area. Such ambulacral plating of the present specimen

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Fig.21. Ophiomusium, or a closely related genus (see Jagt, 2000a), occurs in the Gronsveld, Emael and Meerssen members, preserving (partial) discs with arms attached that reveal all details of disc plating, mouth frame and arm structure; A, NHMM MD 4500; B, NHMM MD 4499 (photographs: Mart Deckers). Disc diameters: 17 and 15 mm, respectively.

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Fig.22. (?)New astropectinid, NHMM MD 5364.12 [greatest diameter: 46 mm]; ENCI-HeidelbergCement Group quarry (Maastricht); uppermost Gronsveld Member (photograph: Mart Deckers). flint eggs must have had a chequered history of Netherlands), the Campine (Kempen) area in repeated erosional and depositional events; they northeast Belgium and into the southwesterly can only be dated (roughly) if they contain age- portion of the German Bundesstaat Nordrhein- diagnostic fossils. It may well be that these Westfalen. flints originate from more southeasterly areas ( occurrences in northwest France and Acknowledgements southern Belgium), having been transported to more easterly regions by (pre-) Pleistocene For donation of study material, close co- rivers (compare Gibbard and Lewin, 2016; operation and helpful discussions, we wish to Garcia-Moreno, 2017; thank a number of museum and university https://www.qpg.geog.cam.ac.uk/research/proj colleagues as well as private collectors, ects/tertiaryrivers/neog.html). Additional amongst whom Bert Boekschoten, Aad van den material is needed, which is why we call upon Engel, Dirk Eysermans, Stijn Goolaerts, Ludo local collectors to focus on such flint eggs in Indeherberge, Henk Jacobs, Paul Kisters, gravel deposits across southern Limburg (the Roland Meuris, Leo Nelissen, Werner Peters,

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Fig.23A, B. Chomataster acules Spencer, 1913, NHMM Van Rijsselt Collection (unregistered); CBR-Romontbos quarry, Eben Emael (Liège, Belgium); upper Nekum Member, Maastricht Formation (photographs: Rudi Dortangs [A]; Maastricht UMC+ [B]). Greatest length of nodule: 178 mm.

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Fig.24A-C. Bathysalenia sp., NHMM 2018 026; internal flint mould, in various aspects (A-C), coated with ammonium chloride prior to photography. Gravel pit ‘De Diepeling’, Tienray (northern Limburg, the Netherlands); erratics from Pleistocene River Maas gravel deposits. Greatest test diameter: 11.6 mm (photographs: Barry van Bakel).

Robert Pieters, Sjir Renkens, Eric van Rijsselt, Haccourt) and ‘t Rooth-Sibelco (Bemelen). For Willy van Rijsselt, Jacques Severijns, Jules pertinent comments on an earlier typescript, we Snellings, and for access to their quarry thank Dr Andreas Kroh (Naturhistorisches grounds, we thank the proprietors of CBR- Museum Wien, Vienna) and Tomasz Borszcz Lixhe (Lixhe) and CBR-Romontbos (Eben (Department of Marine Ecology, Institute of Emael), ENCI-HeidelbergCement Group Oceanology, Polish Academy of Sciences, (Maastricht), Kreco (formerly CPL SA, Sopot).

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Fig.25A-E. Bathysalenia sp., NHMM 2018 026; as Fig. 24A-C, silicone rubber cast of the same specimen, coated with ammonium chloride prior to photography (photographs: Barry van Bakel).

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Fig.26. Cidaroid spines in erratic flint cobble (so-called ‘Maasei’, or ‘Meuse egg’). A, B. Hirudocidaris hirudo (Sorignet, 1850). A, primary spine (greatest length: 29.5 mm, photograph and collection: Werner Peters), beach replenishment at the so-called Zandmotor beach of the North Sea, province of Zuid-Holland, the Netherlands; B. primary spine (NHMM MD 5752, greatest length: 21 mm, photograph: Mart Deckers); C, indeterminate primary spine of cidaroid (NHMM MD 5752) on opposite side of cobble (greatest length: 14.2 mm).

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